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Chapter 27.

Responses of Southeastern

Amphibians and Reptilesto Forest Management: A Review

Kevin R. Russell,T. Bently Wigley, WilliamM. Baughman, Hugh G.Hanlin, and W. Mark Ford1

Abstract—Forest managers in the Southeastincreasingly need information about amphibianand reptile responses to silvicultural practicesin order to guide sustainable forestry programs.A review of existing literature indicates that effectsof silvicultural practices on herpetofauna oftenare region- and species-specific, with individualtaxa responding positively, negatively, or not atall in the short term. Responses of herpetofaunato forestry likely are influenced by adaptationsof taxa to historical disturbance regimes. Fewstudies have evaluated long-term populationor landscape-level implications of silviculturalpractices for herpetofauna. Furthermore, manyexisting studies lack pretreatment data, replication,or appropriate reference conditions. We suggestthat future research focus on manipulative andretrospective studies designed to identify forestrypractices that successfully blend economicobjectives with herpetofaunal conservation.

INTRODUCTION

Forests of the Southeastern United Statessupport a rich diversity of amphibians andreptiles (herpetofauna). Of the more than

450 species of herpetofauna native to NorthAmerica, approximately half occur in theSoutheast and roughly 20 percent, are endemic.Over 100 species (45 amphibians, 59 reptiles,excluding sea turtles) have been reportedfrom the Coastal Plain of South Carolina alone(Zingmark 1978). Herpetofauna often are themost abundant vertebrates in forest ecosystems(Burton and Likens 1975, Congdon and others1986); in the Southeast, they comprise up to 45percent of vertebrate species, excluding fish(Vickers and others 1985).

Several interrelated factors account forthis regional herpetofaunal diversity, includingtremendous variability in habitats related to acomplex matrix of physiography and disturbanceregimes (Sharitz and others 1992). Moreovermany species of southeastern herpetofaunaexhibit biphasic life histories, occupying bothterrestrial and aquatic habitats during annualcycles (Gibbons and Semlitsch 1991).

Increasingly, forest managers are challengedto balance production of forest productswith maintenance of environmental quality,management of wildlife habitat, and conservationof biodiversity (Moore and Allen 1999, Sharitzand others 1992). Concerns about even-agedmanagement, and particularly clearcutting,have prompted considerable research on effectsof timber harvesting on wildlife. Most researchhas focused on mammals and birds, and othervertebrates such as amphibians and reptiles havereceived less attention (deMaynadier and Hunter1995, Gibbons 1988, Moore and Allen 1999).

1 Assistant Professor of Wildlife Ecology andManagement, University of Wisconsin – StevensPoint, College of Natural Resources, Stevens Point,WI 54481; Forest Wildlife Manager, National Councilfor Air and Stream Improvement, Inc., Clemson, SC29634; Southern Region Wildlife Biologist, WestvacoCorporation, Summerville, SC 29484; Professorof Biology, University of South Carolina Aiken,Department of Biology and Geology, Aiken, SC 29801;Research Wildlife Biologist, U.S. Department ofAgriculture Forest Service, Northeastern ResearchStation, Parsons, WV 26287, respectively.

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Despite their presumed role in forest food webs(Burton and Likens 1975, Congdon and others1986), potential value as indicators of habitatquality (Dunson and others 1992), and controversyabout global amphibian declines (e.g., Pechmannand others 1991), herpetofauna often are not fullyconsidered in forest management decisions(deMaynadier and Hunter 1995).

Questions about the compatibility of forestryand conservation of herpetofaunal biodiversity aredriven largely by concerns that both terrestrialand aquatic habitats for many species may bedegraded or eliminated in intensively managedforests. In particular, the permeable eggs, gills,and skin of amphibians make them potentiallysensitive to changes in both aquatic and terrestrialhabitats (Dunson and others 1992). To evaluatethese concerns, deMaynadier and Hunter (1995)presented a comprehensive review of availableliterature about effects of forestry on NorthAmerican amphibians. Several studies suggestedthat clearcutting and other forest managementprescriptions had short-term impacts on someamphibians, especially salamanders. However,other work indicated that many species (1) wererelatively insensitive to forest management,(2) recovered more rapidly after harvesting thanpreviously thought, or (3) responded positivelyto forestry practices (deMaynadier and Hunter1995). This literature review revealed thatamphibian responses to forest managementwere complex and often specific to taxa orregions (deMaynadier and Hunter 1995).

Since deMaynadier and Hunter’s (1995) review,additional studies have provided new insightsabout southeastern forestry and herpetofauna.Also deMaynadier and Hunter’s (1995) reviewdid not address questions about reptiles, perhapsbecause of the focus on global amphibian declines(Gibbons and others 2000), or the historicalperception that forestry impacts on reptilesgenerally were neutral or positive (Campbelland Christman 1982, Welsh and Lind 1991).Although evolutionary, morphological, behavioral,and ecological differences between amphibiansand reptiles are substantial (Gibbons and others2000), it is likely that these ectothermic tetrapodswill continue to be considered collectively fromboth conservation and management perspectives(Gibbons and Stangel 1999, Gibbons and others2000). The purpose of this chapter is to providean up-to-date overview of information availableabout responses of amphibian and reptilepopulations to forestry practices in theSoutheastern United States.

OVERVIEW OF LITERATURE ON FORESTRYAND SOUTHEASTERN HERPETOFAUNA

Harvesting and Silviculture

P resumably the microclimatic, vegetational,and structural changes that occur after timberharvesting, and clearcutting in particular,

create unsuitable conditions for moisture- andtemperature-sensitive amphibians. DeMaynadierand Hunter (1995) reviewed potential negativeeffects of harvesting on microhabitats correlatedwith amphibian species richness and abundance.Timber harvesting removes forest canopy, and socauses increased light penetration that results inhigher soil temperatures and more evaporativeloss of water from the soil and understory.Cover, in the form of leaf litter, coarse woodydebris (CWD), and understory vegetation maybe reduced following clearcutting and associatedsite preparation activities (Hunter 1990). Clearcutareas also are subject to greater daily fluctuationsin temperature and humidity, and to increasedsoil surface disturbance during intensive harvestactivities (deMaynadier and Hunter 1995).However, it has been suggested by several authors(e.g., Campbell and Christman 1982, Greenbergand others 1994, Welsh and Lind 1991) thatclearcutting and other harvesting regimes oftencreate favorable habitats for heliothermic reptilesadapted to early successional habitats.

Amphibians—Several studies in hardwood forestsof the Southern Appalachians appear to supportthe contention that changes in microhabitatsand climate after clearcutting reduce amphibiandiversity and abundance, with negative effectsmost pronounced on salamanders (Ash 1988, 1997;Buhlmann and others 1988; Ford and others 2002;Harpole and Haas 1999; Knapp and others 2003;Petranka and others 1993, 1994). In northernGeorgia, stand age was an important factorexplaining the abundance and communitycomposition of plethodontid salamanders, e.g.,Plethodon and Desmognathus spp., in covehardwood communities (Ford and others 2002).In North Carolina, populations of plethodontidsalamanders in recent clearcuts were 40 percentof those in undisturbed forested plots, and by thefourth year after harvesting, no salamanders couldbe found on clearcut sites (Ash 1988). SimilarlyPetranka and others (1993, 1994) found thatplethodontid salamanders disappeared fromAppalachian forests after clearcutting and thatrecovery to preharvest population levels took up to60 years at high-elevation sites. Hyde and Simons(2001) also reported that effects of disturbance onthe diversity and abundance of plethodontid

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salamanders in the Great Smoky MountainsNational Park were still evident after 60 years.Petranka and others (1993) hypothesized thatduring the last century, clearcutting reducedplethodontid salamander abundance by 70 percentin western North Carolina alone, with currentharvest-related losses approaching 14 millionsalamanders per year.

Three recent studies have evaluated effectsof uneven-aged harvesting techniques onAppalachian salamanders. Harpole and Haas(1999) compared abundance of plethodontidsalamanders before and after application ofseven treatments (understory removal, groupselection, two variants of shelterwood, leavetree, clearcutting, reference) in low-elevationhardwood forests in southwest Virginia. Theyfound that salamander numbers were lower afterharvesting on the group selection, leave tree, andclearcut sites, but no postharvest differences weredetected during the same period on referenceor understory removal sites. However, Fordand others (2000) detected no differences inabundance of plethodontid salamanders amonggroup selection treatments, two-aged timberharvests, and uncut control stands in high-elevation, Southern Appalachian hardwood forestsof North Carolina. Bartman (1998) did not findthat shelterwood harvesting affected salamanderpopulations in the North Carolina Appalachians.

Although it appears likely that diversity andabundance of plethodontid salamanders woulddecrease after clearcutting, Ash and Bruce (1994)and other authors (Ash 1997, Johnson and others1993) argue that available data do not indicate thatthe long-term losses predicted by Petranka andothers (1993, 1994) have occurred. For example,Ash (1997) determined that plethodontidsalamander populations on previously clearcutsites in the mountains of western North Carolinareturned to 100 percent of those in nearbyunharvested forests within 24 years of cutting,rather than the 60 years reported by Petranka andothers (1993). Harper and Guynn (1999) alsoreported that plethodontid salamanders appearedto recover relatively quickly from clearcutting,with salamander densities in stands 13 to 39 yearsold (χ = 21 years) equal to those in older (≥ 40years) stands.

Responses of amphibians to forest managementin other physiographic regions of the Southeastare more complex, with studies reportingindividual species increasing, decreasing, or notchanging in abundance after clearcutting (Clawsonand others 1997, O’Neill 1995, Pais and others

1988, Perison and others 1997, Russell and others2002b). Perison and others (1997) reported thatthe overall abundance of amphibians was lowerin clearcuts than in forested stands in the upperCoastal Plain of South Carolina, but they foundthat certain species, such as green treefrogs (Hylacinerea Schneider) and eastern narrowmouthtoads (Gastrophryne carolinensis Holbrook),were more abundant on clearcut sites. In Alabama,Clawson and others (1997) found that clearcuttingof forested floodplains along blackwater streamshad little impact on the total abundance ofamphibians, but species evenness changed almostimmediately after harvesting. Significant declinesof two-lined salamanders (Eurycea cirrigeraGreen) and gray treefrogs (H. chrysocelis Cope)on clearcut sites were offset by increases of severalspecies, including southern cricket frogs (Acrisgryllus LeConte), southern toads (Bufo terrestrisBonnaterre), and eastern narrowmouth toads.Abundance and richness of several frogs andtoads (anurans) increased at temporary wetlandsin Florida (O’Neill 1995) and South Carolina(Russell and others 2002b) after clearcutting ofsurrounding upland pine plantations. Foley (1994)reported that clearcuts in eastern Texas supportedfewer marbled salamanders (Ambystoma opacumGravenhorst) than did unharvested controls,but timber harvesting had no effect on numbersof smallmouth salamanders (A. texanum Matthes).In a manipulative experiment, Chazal andNiewiarowski (1998) found no significantdifferences in the number of captures, body massand length, or clutch size of pond-breeding molesalamanders (A. talpoideum Holbrook) after 6months exposure to a 4-month-old clearcut and a40-year-old pine stand (animals were captured atan isolated wetland breeding site and placed in100-m2 pens installed after timber harvesting).

Limited evidence suggests that speciescomposition and structure of stands influencediversity and abundance of amphibians in southernforests. Means and others (1996) speculatedthat conversion of natural longleaf pine (Pinuspalustris Mill.) stands to slash pine (P. elliottiiEngelm.) plantations in Florida eliminatedpopulations of flatwoods salamanders (A.cingulatum Cope). In the Coastal Plain of SouthCarolina, Bennett and others (1980) and Hanlinand others (2000) found that the density ofamphibians was significantly higher in natural oak-hickory habitats than in previously clearcut even-aged slash pine plantations. Some researchershave speculated that because habitat featureswhich affect the abundance of amphibians, suchas soil acidity, leaf litter depth and type, hardwood

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shrub abundance, and CWD, may be reducedin conifer plantations, these stand types may beinhospitable for many species of amphibians(Bennett and others 1980, deMaynadier andHunter 1995, Pough and others 1987). However,Hanlin and others (2000) found that pineplantations actually supported higher amphibiandiversity than did hardwood stands. Grant andothers (1994) also reported relatively highamphibian diversity in Coastal Plain pineplantations, with intermediate-aged (8 years old)intensively managed loblolly pine (P. taeda L.)stands having higher amphibian diversity thanrecently clearcut (1 to 3 years old) and olderstands (26 years old). Grant and others (1994)hypothesized that the greater structural andvegetational complexity of intermediate-agedstands, particularly near ground level, couldexplain differences in species diversity. Theysuggested that maintenance of stand structuraldiversity is critical for sustaining herpetofaunalcommunities in managed forests of the Southeast,but this hypothesis remains to be tested.

We believe that taxonomic differences amongamphibians and habitat differences amongphysiographic regions largely explain thedivergence between results of studies conductedin the Southern Appalachians and those conductedelsewhere in the Southeast. Populations ofplethodontid salamanders often decline aftertimber harvesting, but anurans often respondfavorably to harvesting in Coastal Plain forests.Plethodontid salamanders are lungless andentirely terrestrial (Duellman and Trueb 1986) andthese traits may make them sensitive to changes inmicroclimate and microhabitats after harvesting(deMaynadier and Hunter 1995). Results ofstudies from other regions of North Americasupport the supposition that plethodontids mayexperience greater population declines aftertimber harvesting than other groups(deMaynadier and Hunter 1995).

Anurans have higher operating and tolerancetemperatures than do salamanders, and they havethe ability to store and reabsorb large quantitiesof water in the bladder, e.g., 20 to 30 percent ofbody mass (Duellman and Trueb 1986). Thesecharacteristics may explain their tolerance towarmer conditions found in harvested stands(deMaynadier and Hunter 1995). Unlikeplethodontid salamanders inhabiting Appalachianforests shaped by relatively small-scale and low-intensity natural disturbances (Brose and others2001, Sharitz and others 1992), amphibians in thesoutheastern Coastal Plain presumably are

adapted to the high-intensity natural disturbances,e.g., stand-replacing fires, hurricanes, thatcharacterize this region. In much of the CoastalPlain, elevated water tables, increased soilsaturation, and ruts created by tree removal,skidding, and bedding often create standingwater in clearcuts (Cromer and others 2002,O’Neill 1995, Perison and others 1997). Thesefish-free pools, which are often numerous afterheavy rains, apparently attract more anurans toclearcuts than are attracted to unharvested stands(Clawson and others 1997, Cromer and others2002, O’Neill 1995, Perison and others 1997,Russell and others 2002b).

Reptiles—Terrestrial reptiles generally arethought to benefit from the early successionalhabitats created by forest management (Campbelland Christman 1982, Welsh and Lind 1991), butin reality they do not respond to harvesting asa cohesive assemblage. Studies in Florida sandpine [P. clausa (Chapm. ex Engelm.) Vasey exSarg.] -scrub habitats have shown that manyreptile species respond favorably to even-agedforest management (Campbell and Christman1982, Greenberg and others 1994), leading theseauthors to suggest that properly managedclearcutting is sufficiently similar to the effectsof historic high-intensity wildfires so that itsuse can be recommended for maintaining earlysuccessional habitats for reptiles. The integrityof these open-scrub reptile communities isdiminished by forest maturation, and clearcuttingappears to create important microhabitat featuressuch as patches of bare sand (Greenberg andothers 1994).

Although numbers of several lizard speciesincreased following clearcutting in easternTexas, no changes were detected for severalother reptiles (Foley 1994). Clearcutting adjacentto bottomland hardwood stands in the upperCoastal Plain of South Carolina generallyincreased richness and abundance of reptilesrelative to richness and abundance in forestedstands (Perison and others 1997). However, at leasttwo reptile species, ringneck snakes (Diadophispunctatus Linnaeus) and eastern musk turtles(Sternotherus odoratus Latreille), were moreabundant on unharvested plots (Perison and others1997). Seldom encountered in habitats lackingcover, ringneck snakes are among thosesoutheastern reptiles associated with deep litteror other surface objects in mesic hardwood orhardwood-pine forests (Gibbons and Semlitsch1991). Russell and others (2002b) also found thatclearcutting adjacent to Coastal Plain isolated

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wetlands temporarily reduced numbers of severalturtle and snake species, including black racers(Coluber constrictor Linnaeus), but no effectswere evident by 2 years after harvesting. Althoughblack racers are common in early successionalhabitats, clearcutting temporarily eliminated ordisturbed understory vegetation and woody coverthat served as refugia and nest sites.

Although effects of forest management onsoutheastern reptiles have not received the sameattention as those on amphibians, available datasuggest that reptile responses also are species-and region-specific. The response of an individualreptile species to harvesting is influenced by avariety of factors including the degree of habitatspecificity, the spatial scale at which the organismselects its habitat, the morphology and physiologyof the organism, and numerous other biotic andabiotic factors. Thus clearcutting may be sufficientto create the open habitats favored by manysoutheastern reptiles, but insufficient to createhabitat suitable for others unless forested patchesor CWD are retained.

Roads and Skidder RutsMany forestry operations incidentally create

aquatic habitats that are used by herpetofaunafor reproduction, foraging, and cover. Examplesof such habitats include pools along logging roadsand machinery ruts within stands. However, theseactivities can alter hydrological processes anddamage natural aquatic habitats (deMaynadierand Hunter 1995). Because the reproductivestrategies, e.g., timing, of many amphibian speciesare adapted to fluctuating hydrology, an increasingconcern is that these artificial aquatic habitats mayact as population sinks for amphibians if seasonaldrying occurs too rapidly (reproductive failure)or not at all (permanent habitat for predators).To date, only two studies have evaluated effectsof roads and harvest skidder ruts on southeasternherpetofauna. Adam and Lacki (1993) documentedwidespread use of forest road-rut ponds forbreeding by eight species of salamanders andanurans in Kentucky. Road-rut use was positivelyassociated with surface area, depth, and waterclarity, but negatively associated with detritalcoverage. More recently, Cromer and others (2002)compared herpetofaunal communities in recentlyharvested gaps, skidder trails, and undisturbeddepressional wetlands to assess effects of groupselection harvesting and skidder traffic onherpetofauna in a South Carolina bottomlandhardwood forest. Total species richness andabundance were similar among gaps, skiddertrails, and undisturbed bottomland depressions.

However, salamander abundance, especiallyfor pond breeding Ambystoma spp., wasnegatively correlated with pronounced ruttingfrom skidder trails. The characteristic ephemeralhydrology of bottomland depressions was alteredin the harvested gaps and along skidder trails toproduce perennially flooded ponds. This createdpermanent habitat for several aquatic andsemiaquatic species of amphibians and reptilesthat dispersed from bottomland depressionsduring periods of drought. However, the skidder-trail ruts also supported fish and invertebratepredators whose populations in the naturaldepressions typically are controlled byannual droughts.

Although selective harvesting techniqueshave been recommended as an alternative toclearcutting as a means of protecting forestherpetofauna (deMaynadier and Hunter 1995),these approaches may require repeated standentries with additional ground disturbance andmay create more roads and ruts than do even-aged regeneration methods. The artificial aquatichabitats created by these activities may havesignificant implications for habitat selection, andeffects on reproductive success and survival ofherpetofauna should be evaluated.

Site PreparationMechanical treatments—As deMaynadierand Hunter (1995) noted, generalizations aboutthe effects of clearcutting on herpetofaunacan be misleading because a wide range of sitepreparation techniques are associated witheven-aged management. For example, intensivemechanical site preparation is used extensivelyin the Coastal Plain to expose seedbeds andremove competing vegetation prior to replanting,but is rarely employed in Appalachian forestmanagement. Unfortunately, few studies havespecifically examined effects of postharvestmechanical site preparation on southeasternamphibians and reptiles. The available literaturesuggests that these activities can, at leasttemporarily, reduce habitat complexity and affectsome herpetofauna negatively. Although directmortality is likely for selected species (Dodd 1991,Russell and others 2002b), mechanical treatmentstypically are applied only once during standinitiation, and intensive mechanical treatments,such as raking, harrowing, disking, chopping,bedding, probably do greater harm by removingleaf litter, CWD, herbaceous vegetation, rootchannels, and other important microhabitatsfor herpetofauna (Enge and Marion 1986, Whilesand Grubaugh 1993).

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Enge and Marion (1986) comparedherpetofaunal populations of three pine flatwoodsstands in Florida: a 40-year-old pine stand and twoclearcuts receiving minimum (roller-drum chop,bed, plant) and maximum (stump removal, burn,windrow, harrow, bed, plant) site preparationtreatments. After treatment, the maximumsite preparation stand had less leaf litter, CWD,and herbaceous vegetation, and had a greaterpercentage of exposed soil, than did the minimum-treatment or reference stands. Amphibianrichness did not vary significantly among thethree stands, but amphibian abundance was lowerin both site preparation treatment stands than inthe reference stand. Intensive site preparationreduced abundance and richness of most reptilespecies, with the largest impact on fossorialsnakes. The authors attributed lower reptileabundance in the maximum site preparationclearcut to elimination of CWD and other coverobjects that served as refugia and nesting sites.However, intensive site preparation appearedto benefit at least one species, the six-linedracerunner (Cnemidophorus sexlineatusLinnaeus), a lizard that prefers open sandy areas.

A limitation of Enge and Marion’s (1986) studyis that effects of site preparation were not isolatedfrom those of harvesting. Russell and others(2002b) found that when compared to clearcut-onlyand reference stands, mechanical site preparationof sites adjacent to isolated wetlands in the SouthCarolina Coastal Plain did not appear to negativelyinfluence amphibians breeding at the ponds. Infact, bronze frogs (Rana clamitans Latreille)migrated into wetlands from site-preparedstands in higher numbers in the second yearafter treatment. Snakes, including black racers,were less abundant within the first 6 monthsafter treatment, possibly in response to physicaldisturbance of nest sites and reductions in groundcover. These effects were short lived, however, andno effects of site preparation on reptiles weredetected in the second year after application.

In addition to removing surface cover,mechanical site preparation may destroy burrowsand other subsurface refugia of fossorialherpetofauna. Several studies have documenteddestruction of gopher tortoise (Gopheruspolyphemus Daudin) burrows by chopping andother mechanical treatments (Diemer and Moler1982, Landers and Buckner 1981, Marshall andothers 1992, Tanner and Terry 1981), althoughLanders and Buckner (1981) and Diemer andMoler (1982) observed tortoises emerging from

destroyed burrows and either reopening themor excavating new sites. Loss of gopher tortoiseburrows to site preparation can indirectly affectother species; at least 332 wildlife species areknown to use burrows of gopher tortoises,including several rare amphibians and reptiles(Lips 1991). Soil disturbance from site preparationalso has been linked to destruction of Red Hillssalamander (Phaeognathus hubrichti Highton)burrows in Alabama (Dodd 1991).

Prescribed fire—Prescribed burning is usedto achieve a variety of silvicultural objectivesincluding controlling heavy fuel accumulation,exposing mineral soil, releasing available nutrientsfor seedbed preparation, and controlling insects,diseases, and competing vegetation. A detailedliterature review of fire effects (and fire exclusion)on southeastern herpetofauna was conductedby Russell and others (1999) and only a briefsummary is provided here. Generally, replacingfire-adapted vegetation with fire-intolerantassociations, e.g., hardwoods, in the southeasternCoastal Plain leads to concomitant declines inoverall herpetofaunal abundance and diversity.However, it may be appropriate to use prescribedfire in combination with other forestry practicesto benefit Coastal Plain herpetofauna by restoringan historic mosaic of successional stages, habitatstructures, and plant species compositions inboth terrestrial and aquatic habitats (citationsin Russell and others 1999). For example, insouthern Florida, richness and abundance ofherpetofauna consistently were higher in slashpine plots subjected to three different burnintervals (1, 2, 7 years) than in a reference plotprotected from burning for 20 years (Mushinsky1985). Based on these results, Mushinsky (1985)recommended a 5- to 7-year prescribed burn cycleto maintain diverse herpetofaunal communities insouthern Florida sandhills.

Available evidence suggests that directmortality of herpetofauna following fire typicallyis low and presumably outweighed by maintainingdesired habitat features (Means and Campbell1981, Russell and others 1999). Although fire-induced disturbance may temporarily decreaseherpetofaunal diversity within a particular stand,a heterogeneous matrix of stand ages andstructural conditions should increase diversityon a broader scale (Greenberg 2002, Greenbergand others 1994, Jones and others 2000, Litt andothers 2001). Unfortunately, concerns over croptree productivity, smoke management, air qualitystandards, and liability have led to fire exclusion

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policies that may have significant long-termconsequences for herpetofauna in Coastal Plainforests and elsewhere (Russell and others 1999).

Even within fire-adapted southern forests somespecies of herpetofauna may depend on climaxvegetation (Greenberg 2002). Means and Campbell(1981) examined herpetofaunal communities inlongleaf pine and shortleaf pine (P. echinata Mill.)stands in peninsular Florida that had beenburned annually for 60 to 70 years and in anunburned forest that had succeeded to a closed-canopy hardwood association. Three species ofamphibians [tiger salamander (A. tigrinumnebulosum Holowell), oak toad (Bufo quercicusHolbrook), ornate chorus frog (Pseudacris ornataHolbrook)] and six-lined racerunners werecaptured predominantly from the burnedpine stands, whereas three amphibian species[marbled salamander, mole salamander, and slimysalamander (Plethodon glutinosus Green)] werecaptured almost exclusively in the hardwoodforest. The authors suggested that thesedifferences in distribution reflected adaptations(or lack thereof) of individual species to fire(Means and Campbell 1981).

Almost all studies of fire effects on southeasternherpetofauna have been conducted in Coastal Plainforests (Russell and others 1999), and caution mustbe exercised when extending conclusions to otherareas. Until recently, fire was not considered animportant or desirable disturbance regime inmixed-hardwood forests of the Appalachianand Piedmont regions (Brose and others 2001).However, it has been hypothesized that periodic,low-intensity surface fires were crucial forperpetuating these oak-dominated forests formillennia and are necessary to restore such forests(Brose and others 2001). To date, only two studieshave investigated prescribed fire-herpetofaunarelationships in these areas. Ford and others(1999) found that prescribed fires in the SouthernAppalachians had little effect on herpetofauna andconcluded that concerns about negative effects ofprescribed burning on plethodontid salamandersprobably were unwarranted. An ongoing studyevaluating the use of prescribed fire to restore oakforests in the South Carolina Piedmont also hasnot found dramatic negative impacts (Floyd andothers 2002).

Other topics needing attention include (1)the combined effects of fire frequency, intensity,and seasonality on herpetofauna; (2) the use ofherbicides as a substitute for prescribed fire(Litt and others 2001); and (3) the use of

prescribed fire to restore and maintain aquatichabitats of herpetofauna threatened by hardwoodsuccession (Russell and others 1999).

Herbicides—In forestry, herbicides are used forsite preparation, for release of crop trees fromherbaceous and woody plants, for managingspecies composition and structure, and for timberstand improvement (Miller and Mitchell 1994).Herbicides may be broadcast across a stand,sprayed in bands centered on rows of trees,or applied to individual woody stems. Individualstems usually are treated by directly sprayingfoliage, applying the herbicide to the tree bole (orto wounds on the bole), or applying a soil-activeherbicide to the ground near the tree.

Documented adverse effects of herbicideson some herpetofaunal life stages includemortality, reduced body mass, failure tometamorphose, decreased stimulatory responseof neuroepithelial synapses, chromosomalfragmentation, deformities, and DNA profileabnormalities (citations in Pauli and others 2000).It has been suggested that herbicides are amongthe causative factors explaining global declinesof amphibian populations (citations in Fellersand others 2001). However, these effects generallyhave occurred at exposure levels above thoselikely to occur in normal forestry operations.Furthermore, several literature reviews haveconcluded that commonly used forestry herbicidesare not acutely toxic to wildlife because they haverelatively low mutagenicity, have no or very weakoncogenetic effects, are rapidly eliminated byanimals, do not bioaccumulate, and have a shortenvironmental half life (McComb and Hurst1987, Miller and Witt 1991). Forestry herbicidesalso are used infrequently, i.e., many even-agedstands receive only one or two applications duringa typical rotation, and most herpetofauna likelyare shielded from direct exposure, i.e., by beingunderground or under vegetation, leaf litter,or CWD.

Because herbicides are designed to killvegetation, they can affect herpetofauna indirectlyby altering habitat. Herbicide effects on habitatvary with soils, structure of the pretreatmentplant community, herbicide product used,application rates, timing of application, weatherconditions, and other factors. However, herbicideapplication to individual trees in midrotation ormaturing stands often promotes canopy gaps andunderstory biomass production (McComb andHurst 1987). When broadcast in regeneratingstands, herbicides often temporarily reducebiomass production for one to several growing

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seasons, and shift the dominant understoryvegetation from woody to herbaceous plants(Miller and Witt 1991).

Few studies have documented herpetofaunalresponse to herbicide-induced habitat changes.Results of those studies, and studies for otherwildlife taxa, suggest that herpetofaunal responsesare species-specific (Howell and others 1996,Lautenschlager 1993, McComb and Hurst 1987),with individual species increasing, decreasing,or not changing in abundance at the stand level(Cole and others 1997, Harpole and Haas 1999,Lautenschlager and others 1998, Yahner andothers 2001). Landscape-level responses ofherpetofaunal species to herbicide applicationsprobably depend on the productivity and naturaldisturbance regime of the landscape (Huston1999), the extent of the area simultaneouslytreated with herbicides, the vegetation structureof treated stands and the broader landscape,and other factors previously described.

Whether used alone or with other managementpractices, e.g., prescribed fire, herbicides may beapplied to meet selected management objectivesfor herpetofauna and other wildlife species. Forexample, Brooks and others (1993) concludedthat any of the three herbicide treatments theyevaluated (hexazinone, imazapyr, and picloram+ triclopyr) were compatible with the goal ofmaintaining quality habitat for gopher tortoises.Managers can use herbicides to control nonnativeplant species; create snags; manipulate the speciescomposition and structure of understory, midstory,and overstory vegetation; manage the spatial andtemporal availability of habitat; and for otherpurposes (Wigley and others 2002).

Riparian Buffers, Isolated Wetlands,and Terrestrial CorridorsRiparian buffers—Retention of streamsidemanagement zones (SMZs or buffers) as a meansof conserving biodiversity continues to be a widelydebated strategy (Harrison and Voller 1998). Somestudies conducted in the Pacific Northwest suggestthat unharvested riparian buffers are importantfor protecting stream- and riparian-associatedamphibians from effects of timber harvesting(Corn and Bury 1989, Welsh and Lind 1991).Riparian buffers presumably lessen accumulationof fine sediments in stream substrates, limitincreases in water temperatures, and mitigateother negative impacts of soil transport and solarradiation on stream habitats (deMaynadier andHunter 1995). Little information is available,however, about effects of riparian logging on

southeastern stream amphibian communities(Pauley and others 2000). In the SouthernAppalachians, salamanders were 50 percentmore abundant in SMZs than in adjacentharvested areas (Petranka and others 1993).In the western Piedmont of North Carolina,Willson and Dorcas (2003) found that the relativeabundance of stream-dwelling salamanders wasinversely proportional to the percentage ofdisturbed habitat at the watershed scale, butthey found no relationship between the relativeabundance of salamanders and the percentageof disturbed habitat within riparian buffer zones.Stiven and Bruce (1988) speculated that stream-dwelling blackbelly salamanders (Desmognathusquadramaculatus Holbrook) were less abundantin recently logged Appalachian watersheds, andthat harvesting also might alter genetic diversityof the affected populations.

In eastern Texas, Foley (1994) found that SMZsretained in clearcuts actually supported higherdiversity of herpetofauna than did unharvestedreference stands. He and others (deMaynadierand Hunter 1995) have suggested that in additionto protecting aquatic amphibians, riparian bufferstrips could also provide an intact strip of forestedhabitat capable of harboring populations for futurerecolonization of adjacent disturbed areas. Bowersand others (2000) examined herpetofaunalresponse to different planting regimes in the PenBranch corridor, which is associated with a third-order stream on the Savannah River Site in SouthCarolina. This stream received thermal effluentsfrom a nuclear reactor for over 30 years, andthis resulted in the destruction of most riparianvegetation in a portion of the stream’s floodplain.Subsequent erosion created a braided streamsystem with a greatly expanded delta, andrestoration of the area began with planting ofbottomland hardwood species in 1993. Speciesdiversity of herpetofauna in the unaffectedriparian zone was significantly higher thanon vegetated islands located between streambraids within the impacted floodplain corridor,and there were also significantly more speciesand individuals within the riparian zone thanin the corridor. According to Bowers and others(2000), these results highlight the importance ofthe unaffected riparian zone in the faunal recoveryof the floodplain.

Recommended streamside buffer widthsfor herpetofauna in other regions of NorthAmerica range from 30 to over 100 m (McComband others 1993, Rudolph and Dickson 1990).

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It has been recommended that riparian bufferwidths be adjusted proportionally with streamwidth, intensity of adjacent harvest, and slope(deMaynadier and Hunter 1995). However,we agree with Wigley and Melchiors (1993)that we know too little about empiricalrelationships between forest managementeffects and riparian habitat functions to justifyour recommending specific stream buffer widthsfor southeastern herpetofauna.

Isolated wetlands—Although little effort hasbeen devoted to research and management ofstream-associated herpetofauna in southernforests, protection of isolated wetland habitatsin the southeastern Coastal Plain has receivedincreasing attention. Carolina bays, cypress ponds,and other isolated wetlands, i.e., those with nopermanent connections to aboveground streamor river systems, are critical habitats forherpetofauna adapted to seasonal hydroperiodsand the absence of predatory fish. Of 29 anuranspecies native to the southeastern Coastal Plain,20 breed primarily or exclusively in isolatedwetlands (Moler and Franz 1987). Several speciesof salamanders, e.g., Ambystoma spp., alsomigrate to isolated wetlands for mating and eggdeposition but return to upland habitats for theremainder of the year (Gibbons and Semlitsch1991). In contrast, many Coastal Plain turtles andsnakes seek food and cover in isolated wetlands ortheir peripheries but migrate to adjacent uplandsfor egg laying and hibernation (Gibbons andSemlitsch 1991, Russell and Hanlin 1999).

Most species of herpetofauna associated withisolated wetlands in the Coastal Plain also useadjacent upland forests, and several authorshave recommended, on the basis of anecdotal orretrospective data, that closed-canopy forestedbuffers or complete exclusion of upland forestmanagement activity is necessary to protectthese aquatic habitats and maintain landscapeconnectivity among wetlands (see citationsin Russell and others 2002b). For example,Pechmann and others (1991) speculated that theinitial absence and then presence of marbledsalamanders at an isolated wetland in SouthCarolina resulted from regeneration ofsurrounding upland forests that previously wereclearcut and burned. Raymond and Hardy (1991)reported that a clearcut 156 m away from abreeding pond in Louisiana appeared to influencethe migratory movements and survivorship of thepond’s breeding population of mole salamanders.On the strength of data on movements of several

salamander species from isolated wetlandsto adjacent upland forests, Semlitsch (1998)hypothesized that a buffer zone encompassing95 percent of the populations using those uplandstands would extend approximately 164 m fromthe wetland’s edge. Burke and Gibbons (1995)estimated that an upland buffer 275 m in widthwould be necessary to protect 100 percent of thenest and hibernation sites of two aquatic turtlespecies associated with isolated wetlands.

In contrast, Wigley (1999) reported thatretention of an adjacent forested buffer wascorrelated with the presence of only 1 of 40amphibian species and 37 reptile species sampledfrom 444 temporary isolated wetlands acrossthe southeastern Coastal Plain—the pine woodstreefrog (Hyla femoralis Bosc). Russell and others(2002a) also found that 5 small isolated wetlands(0.38 to 1.06 ha) surrounded by 18- to 25-year-oldloblolly pine plantations in the Coastal Plain ofSouth Carolina were used by at least 56 speciesof herpetofauna, suggesting that these aquatichabitats within managed forests are capable ofsupporting high herpetofaunal diversity. Althoughretaining forested buffers around isolatedwetlands is widely recommended, to date onlyRussell and others (2002b) have experimentallyevaluated management of upland forest bufferson southeastern wetland herpetofauna. Theyexamined immigration and emigration ofherpetofauna from isolated wetlands in theSouth Carolina Coastal Plain before and afterclearcutting and mechanical site preparationof adjacent upland forests. Although harvesttreatments significantly altered overstory andground cover characteristics of upland stands, notreatment-related changes in the overall richness,abundance, or community similarity of amphibianand reptile communities at the wetlands wereobserved. Only short-term negative effects wereobserved for turtles and snakes. These taxa wereless abundant only within the first 6 months afterclearcutting and site preparation, possibly inresponse to physical disturbance of nest sites andtemporary changes in ground cover. No amphibianspecies showed negative responses to treatments,and the number of bronze frogs at the wetlandsincreased after treatments. The authors noted thatalthough it is premature to suggest that uplandforested buffers surrounding southern isolatedwetlands are unnecessary, assumptions abouteffects of forestry operations on isolated wetlandherpetofauna, and management based on suchassumptions (Semlitsch 1998, 2000), mustbe tested in the field.

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Corridors—Preston (1962) was among the firstto suggest possible conservation benefits of uplandhabitat corridors. Preston speculated that habitatpreserves would become isolated, and that the onlyremedy was to maintain continuous corridors thatwould link reserves. Most studies of corridors haveexamined movement patterns of mammals andbirds (Bennett 1990, Wegner and Merriam 1979).The function and conservation value of uplandcorridors is still debated widely (citations inHarrison and Voller 1998, Ford and others 2000).Although use of corridors to manage amphibianshas been advocated (Semlitsch 2000), we are awareof only one study of the effects of retaining uplandforest corridors for herpetofauna in the Southeastor elsewhere (Baughman 2000). In this study fromSouth Carolina, three sites were randomlyselected for retention of a 100-m wide unharvestedforest corridor traversing the length of a clearcut,and one site was assigned as an unharvestedreference. Baughman (2000) found that meannumbers of herpetofauna captured entering orwithin corridors did not differ from mean numbersof herpetofauna captured in harvested areas, andthat herpetofauna assemblages and movementrates for corridors were similar to those for thestands from which the corridors were created.Although corridors provided a continuous web ofclosed-canopy forest across the study landscape,Baughman (2000) emphasized that long-termmonitoring is needed before potential benefits ofterrestrial corridors for herpetofauna in managedforests of the Southeast can be determined.

Demographic Responses to ManagementAlthough short-term measures of richness

and abundance may not be affected by forestmanagement, such measures often are not goodpredictors of habitat quality (Van Horne 1983),and changes in habitats could have longer termconsequences for reproductive success, survival,and dispersal of herpetofauna. Few studies havecollected demographic data to determine whetherresponses to forestry practices are age- or sex-specific. Enge and Marion (1986) reported thatalthough there was no difference between overallfrog biomass in forested plots, in clearcut plots,fewer juvenile frogs were captured on harvestedsites. Raymond and Hardy (1991) suggested thatsurvival of female mole salamanders was lowerthan survival of males following clearcutting,whereas Ash (1988) reported that sex and ageclasses of plethodontid salamanders declined atthe same rate after clearcutting. Also, withoutappropriate marking and recapturing techniques,it is difficult to collect data indicative of true

population sizes or to monitor movements ofherpetofauna in response to forestry practices(Ash and Bruce 1994). For example, estimatesby Petranka and others (1993) of plethodontidsalamander mortality resulting from clearcuttingare based on the assumption that thesesalamanders exhibit poor dispersal capabilities and strong site fidelity. Bartman (1998) did notobserve dispersal of plethodontid salamandersimmediately after logging in North CarolinaAppalachian forests, but fates (death vs. dispersal)of herpetofauna after clearcutting and otherforestry activities remain poorly known (Ash andBruce 1994, deMaynadier and Hunter 1995).

Landscape-Level ResponsesAlthough characterizing stand-level

responses of herpetofauna to forest managementis important, perhaps the most pressing questionsare at larger scales (Guerry and Hunter 2002).Some recent studies have characterizedherpetofaunal communities at the landscapelevel. Leiden and others (1999) conducteda broad survey of herpetofauna across anindustry-managed landscape in South Carolina.The landscape contained stands in various standstructural classes, including pine plantations.Leiden and others (1999) confirmed the presenceof 73 of 102 species of amphibians and reptilespotentially occurring in the landscape (basedon range maps). This represented the highestrecorded richness of amphibians and reptilesin South Carolina, with the exception ofthe Savannah River Site, where continuoussampling has occurred since the 1950s (Gibbonsand others 1997).

Responses of herpetofauna to forestfragmentation have not been studied as oftenas have responses of other vertebrates, such asbirds. However, a limited number of field studiessuggest that isolation of forest patches mayinfluence occupancy of terrestrial habitat in suchpatches by adult amphibians (citations in Guerryand Hunter 2002). Fox and others (2004) andShipman and others (2004) censused amphibiansand reptiles in four forested watersheds (1500 to4000 ha each) in the Ouachita Mountains that weremanaged at different intensities, and thus levels of“fragmentation,” ranging from largely unmanagedto intensive even-aged management. Watershed-to-watershed differences in amphibian richnesswere negligible, and community similarities werehigh (Fox and others 2004). The watershedshad similar reptile communities, but the leastintensively managed watershed had lower

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per-plot abundance, species richness, anddiversity of reptiles than the others (Shipman andothers 2004). This was attributed to dominance bytwo reptile species in the least intensivelymanaged watershed.

Because many aquatic and semiaquaticherpetofauna use adjacent terrestrial habitats fordispersal, foraging, and refuge, both the proximityof wetlands to terrestrial habitat and the area ofterrestrial habitat may influence habitatoccupancy. If populations of wetland-associatedherpetofauna exhibit metapopulation structure,reduced immigration and emigration ratesresulting from disconnection of habitat patchesmay negatively influence viability (Guerry andHunter 2002, Joyal and others 2001). In Maine,Guerry and Hunter (2002) found species-specificresponses of pond-breeding amphibians to areaand proximity of adjacent terrestrial forests.Although the presence and abundance of somespecies were positively related to forest area andpond-forest adjacency, other species exhibitednegative or no associations with one or both ofthese factors. However, we are unaware of anystudies that explicitly evaluate effects of forestfragmentation on either terrestrial or aquaticamphibians in the Southeast.

MANAGEMENT IMPLICATIONS OFNATURAL DISTURBANCE REGIMES

Currently available evidence suggests thatsoutheastern herpetofauna respond in acomplex manner to changes in climatic,

vegetational, and structural features of stands andlandscapes after the implementation or exclusionof specific management practices (such as firesuppression). DeMaynadier and Hunter (1995)argue that herpetofauna generally benefit whenforest management prescriptions retain sufficientmicrohabitat and microclimate elements withinstands, and ensure a diversity of habitat typesacross larger areas. They also suggest thatidentifying and then minimizing differencesbetween forest management practices andhistoric patterns of natural disturbance, e.g.,retention or creation of microhabitats, willimprove conservation of herpetofauna in ourmanaged forests.

We suggest this historic context has often beenoverlooked by those considering the effects offorest management on southeastern herpetofauna.Current forest management regimes are only thelatest in a continuum of forest clearing, intensiveagriculture, prescribed burning, forest regrowth,and timber harvesting across the Southeast

(Sharitz and others 1992). Prior to humaninfluence, natural disturbances, e.g., fire,hurricanes, windthrow, ice storms, occurring atdifferent frequencies, intensities, and extentsdepending on physiographic region, controlled thecharacter of southern forests and maintained thestand and landscape diversity essential to supportthe flora and fauna of the region (Brose and others2001, Myers and Van Lear 1998, Sharitz and others1992). Unmanaged southern forests were not ahomogeneous blanket of “intact” or “continuous”closed-canopy forest, but rather a heterogeneousmixture of stands of different ages and structuraltypes. Many vertebrates in the South, includingherpetofauna, have tolerated and adapted todisturbance events throughout much of theirevolutionary histories (Campbell and Christman1982, Greenberg and others 1994, Russell andothers 1999). Thus the complexity and regionalnature of herpetofaunal responses to forestmanagement should not be surprising. As Meansand Campbell (1981) point out, it is illogicalto conclude that herpetofauna associated withsoutheastern forests are not themselves adaptedto local patterns of disturbance. We have found,however, that few studies or managementrecommendations (Semlitsch 2000) involvingresponses of herpetofauna to forest managementhave fully considered the spatial and temporalcomplexity of forest habitats, includingdisturbance scales and intensities that speciesand communities are adapted to. It is absolutelynecessary that we understand this context ifwe are to predict how southeastern herpetofaunawill respond to forest management, and ifwe are to develop efficacious and cost-effectiveconservation strategies. For example, arerecommendations for closed-canopy buffersaround isolated wetlands consistent with theexistence of the exposed and sparsely vegetatednest sites selected by many turtle species (Kolbeand Janzen 2002)? Management and recoverystrategies for herpetofauna that do not recognizethe dynamic rather than static nature of southernforests, or those that provide one-size-fits-allsolutions, are likely to fail.

CONSIDERATIONS FOR FUTURE RESEARCH

Increasingly, researchers and resourcemanagers are recognizing the importanceof herpetofauna within the context of forest

management (deMaynadier and Hunter 1995,Dunson and others 1992). However, much remainsto be learned concerning effects of forestrypractices on southeastern herpetofauna. Currentlyavailable data suggest that herpetofauna are

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influenced both positively and negatively (andoccasionally not at all) by management of southernforests, and responses are specific to individualregions, taxa, and management prescriptions.The population and community effects of forestmanagement activities on southeasternherpetofauna are still difficult to assess, though,because of methodological limitations and becausea variety of study designs have been employed(deMaynadier and Hunter 1995).

The absence of pretreatment data, replication,and true reference conditions in many studieshas limited conclusions about impacts of forestryon herpetofauna (Ash and Pollock 1999,deMaynadier and Hunter 1995, Petranka 1999,Russell and others 1999). Most studies haveinferred management effects on the strengthof retrospective comparisons of herpetofaunalattributes of harvested and unharvested sites.This approach assumes that the herpetofaunalpopulations of harvested sites once exhibitedcharacteristics, e.g., abundance, identical withthose of populations present on forested referencesites. Baseline data on habitat parameters arenecessary if we are to assess the comparabilityof sites and the extent of postharvest changes.Only six studies investigating effects of forestmanagement on southeastern herpetofauna haveemployed manipulative designs with pretreatmentand posttreatment data, treatment replication,or true spatial and temporal references (Ash1997, Chazal and Niewiarowski 1998, Clawsonand others 1997, Harpole and Haas 1999, Knappand others 2003, Russell and others 2002b). Alsoneeded are longer-term studies that separateimmediate population responses to harvestingfrom long-term effects on fitness.

The challenge for future studies ofherpetofauna-forestry relationships has movedbeyond simply documenting the range of harvesteffects to successfully blending economic andcultural objectives with those for conservationof herpetofauna by identifying silviculturalprescriptions that retain significant naturalcomponents of regenerating stands (deMaynadierand Hunter 1995, Grant and others 1994).Although documentation of the magnitude ofsilvicultural effects on herpetofauna is increasing,the causal factors that shape the distribution andabundance of herpetofauna in southern forestsremain poorly understood. Pioneering work byMacArthur and MacArthur (1961) demonstratedthe importance of vegetational structural diversityto avian communities. However, quantitativestudies that explicitly examine relationships

among structural attributes of forests andherpetofaunal populations in the SoutheasternUnited States are lacking (Grant and others1994). Studies from the Pacific Northwest andNortheast suggest that structural characteristicsand components of forests, particularly those atground level, e.g., CWD, leaf litter depth andmoisture, understory vegetation, are importantcorrelates of herpetofaunal abundance anddiversity (Aubry 2000, deMaynadier and Hunter1995, McComb and others 1993, Pough andothers 1987).

Although microhabitat variables such asCWD and leaf-litter depth often increase withstand age, there is a great deal of stand-specificvariability related to natural and silviculturaldisturbance history, climate, soils, elevation,proximity to aquatic habitats, and other influences(Oliver and Larson 1996). For example, intensivesite preparation treatments, e.g., bedding orwindrowing, may retard development of standstructure by eliminating cull trees, snags, CWD,and understory species, whereas less-intensiveapplications that only slightly disturb the soil, e.g.,roller chopping, or occasional prescribed burning,may increase diversity and biomass of understoryspecies (Hunter 1990). Thus stand age may notbe an accurate delimiter of transitions in standstructural development (deMaynadier andHunter 1995, Hunter 1990, Oliver and Larson1996), particularly across regions and ownershipswith different methods of harvesting orsite preparation.

Approximately 90 percent of southeasternforests are privately owned (U.S. Departmentof Agriculture, Forest Service 1988), and mostof these forests will continue to be managedfor economic benefit. We think that informationobtained by means of retrospective andmanipulative studies that elucidate relationshipsamong stand structural diversity, forestmanagement practices, and herpetofaunalcommunities can be used to integrate managementof these forests with the protection and promotionof herpetofaunal biodiversity. This will beaccomplished by approximating the rangeof natural disturbance events that historicallyshaped the region’s forests. One researchapproach is to inventory the distribution andabundance of herpetofauna in forest standswith variable structural characteristics andmanagement histories within larger landscapes(Gibbons and others 1997, Wigley andothers 2000). Then quantitative models can bedeveloped that relate distribution, abundance,

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and demographic characteristics of species tospecific habitat elements found in managedforests, and eventually integrated into sustainablelandscape models that would predict herpetofaunaresponses to different management scenarios(Wigley and others 2001).

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